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The Influence of Weather on Rodent Activity

Article  in  Journal of Mammalogy · February 1981

DOI: 10.2307/1380484

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 THE INFLUENCE OF WEATHER ON RODENT ACTIVITY

W. L. VICKERY AND J. R. BIDER

AnSTRACT.-\Ve measured the activity of PeromysclIs manicll/atlls, Clethriollomys

Mapperi, and Napaeozapus insi{mis llsing a sand transect in a forest in Quebec. Activity
on each of 651 summer nights over 8 years was compared to the weather of each night.
All three rodents were most active on rainy and on warm nights. C. gapped and N.
insignis showed increased activity on cloudy and moonless nights and when rain had
Tlot fallen for several nights. C. gapperi also increased activity with increased humidity.
Variation in activity due to weather was slight and more likely associated with predator
avoidance than with physiological restrictions.

Small nocturnal rodents face a problem in temperature regulation in cold environ-

ments because of their large surface to volume ratio and associated higher metabolic
rate (per g body weight) than larger mammals (Schmidt-Nielsen, 1975). As the differ-
ence between body temperature and ambient temperature (below the thermal neutral
zone) increases, metabolic rate must increase to compensate for increased heat loss.
Thus, one way in which small nocturnal rodents might conserve energy is by restrict-
ing their activity to those times when temperatures are suitable. This should be true
regardless of physiological adaptations for thermoregulation as long as ambient tem-
peratures are below the thermal neutral zone.
Water loss may also be a problem for animals with a high surface to volume ratio.
If so, one might expect them to be active only in moist or humid areas or more active
on wet and humid nights. Falls (1968) has reviewed work on Peromyscus activity and
concluded that although numerous laboratory studies showed a relationship between
activity and temperature and humidity, field studies showed little indication of these
effects. Getz (1968) found winter activity of C. gapped to be independent of temper-
ature and humidity but influenced by cloud cover and rainfall. Brower and Cade
(1966) found N. insignis to be most trappable on cold nights.
In the field, small rodents must solve not only the problems of heat and water
balance but also problems related to food gathering, predator avoidance, and social
interactions. Because small rodents must forage regularly or risk starvation some of
their activity must be independent of weather. However, one would expect the
amount of activity on a given night to be inHuenced hy weather. The extent to which
this is not true is a measure of the independence of small rodents from meteorological
conditions.
We examined the influence of weather on the summer activity of Perom!lsclls
maniculatus, Clethrionomus gapperi, and Na/JaeOz,(/pllS insigl1is, based on data ob-
tained during eight summers. These rodents are known to he influenced by weather
(Falls, 195.3; Brower and Cade, 1966; Bider, 1968; Getz, 1968; Marten, 197,3). How-
ever, the extent to which summer activity is inHuenced by weather has been estimated
only by Marten (1973) based on j\lst one year (19G9) of our data.
METHODS
Rodent activity was measured on a 91.5-m sand transed (Bider, 19(8) starting at a forest edge
and running perpendicular to the edge into the forest. The transect was 45 em wide and covered
hya L'5-m wide polythene canopy ahout 1.5 m ahove the ground. This prevcnted rainfall from
obliterating lra<:ks on the transect.
The forest, mostly maple and hirch with hal sam fir and white spruce in wct arcas, had ,l closed
eanopy and was located .3 km N Lac Carre, Terrehonlle Co., P.Q., Canada (4GomYN, 74°29'W).
The site, topography, climate, and vegetation were descrihed hy Bieler (1968).
The numher of crossings of thc transect hy each species on each night was I1scd as an inde)(

j. Mamm., 62(J);140-145, 1\;)81 140

Febma r u 1981 VICKERY AND BIDER-WEATHER AND RODENT ACTIVITY 141

of the activity of that species on that night. r.,'lost (over 95%) activity by these rodents was
nocturnal. Nightly signs of activity of each species increased through the summer as population
sizes increased. Thlls, activity totals at different times of year were not comparable. For this
reason each night's activity was divided by the mean activity during that part of the slimmer
following the method of DOllcet and Bider (1974) and Vickery and Bider (1978). This transfor-
mation produced normally-distributed aetivity variables.
The amllysis is based on 651 nights between 1 June and f:i September, 1964 to 1971. On other
nights heavy rainfall caused flooding of the transect so that data are not available.
Weather data were collected every 2 h at a Stevenson screen near the tnmsect. Weather vari-
ables measured were: presence of rainfall; time of rainfall [none, morning (0700-1200 h), after-
noon (1.300-1800 h), evening (1900-0000 h) and late night (0100-0600 h)]; amount of rainfall
(total, noon-noon); mean temperature (in shade, 2.5 em above ground) during the evening, late
night, and whole night (1900-0f:iOO h); maximum temperature (daytime); minimum temperature
(nocturnal); temperature at sundown; temperature drop within 2 h at sundown; noeturnal, eve-
ning, and late night cloud cover (estimated as percentage cover and summed over each time
interval); nocturnal (1900-0600 h), evening, late night, and daytime (0700-1800 h) mean relative
humidity; mean barometric pressure; total change in harometric pressure during the day; mean
daily wind speed; wind direction (measured as the numher of hours during which the wind
hlew from the east, a sign of an impending storm); number of days since the Inst rainfall; and
the phase of the moon (on a scale of 0-15; new to full, respectively).
These weather variables were highly correlated with each other. Principal component analysis
was used to produce linear combinations of those weather variables that were independent from
each other and accounted for the maximum amount of variation in weather data. The scores
produced by these principal components were compared with rodent activity using correlation
analysis. This allowed LIS to assess the effects of weather variables independently. Canonical
correlation analysis was used to assess the extent to which all rodent activity varied with weather.

RESULTS
During the study PerolllyscllS /1Ial1iculatlls crossed the transect 10,324 times, Cleth-
rionomys gap peri crossed 4,407 times, and Napaeoza!Jus insigllis crossed 34,953
times.
Principal component analysis reduced the number of weather variables from 23 to
6. These six variables accounted for 76% (31, 15.5, 13.5,6,5, and 5%) of the variation
in the original weather data. The other 17 principal components had eigenvalues less
than one and were not used in the analysis. The standardized hwtor coefficient matrix
showed that the six weather factors can be clearly labelled as: 1) temperature; 2)
relative humidity; 3) cloud cover; 4) rainfall; 5) wind direction, barometric pressure,
and moonlight; 6) moonlight and recent rainfall (Table 1).
Table 2 shows the correlations between rodent activity and weather. All three ro-
dents were most active when rain fell and when temperatures were high. C. gapperi
and N. illsignis increased activity on clolldy nights and moonless nights when rain
had not fallen recently. C. gapperi was also more active when relative humidity was
high. However, neithcr P. IIl(1lliclIlatlls nor N. illsigllis reacted to changes in relative
humidity and no species had activity correlated with the fifth principal component.
Two significant canonical variables explain the relationship between weather and
rodent activity (Tahle 3). The first canonical variable shows that all three rodents react
similarly to weather. That is, they are most adive on rainy, cloudy, warm, moonless,
and highly humid nights and when it has not mined for some time. The second
canonical variahle shows that C. g(lpperi reads to weather differently than the other
two rodents. It is morc dependent on high humidity and rainf~dl than are P. IIWlli(.'lI-
lat!ls and N. illsigllis.
Weathcr explained 18.0% oral! rodent aetivity. By species, weather explained 10.3%
for N. iHSigHis, 8.3% for C. gapperi, and 1.7% f{Jr P. mallicfllatHs. The proportion of
variation ill activity explained hy each wcather flctor for each rodent was estimated
142 JOURNAL OF MAMMALOGY Vol. 62, No.1

TABLE L-Stmuiardized factor coefficient matrix/or the first six principal components of the
weatflcr data olld colltllltHlalilies of the factors wilh the original weather variahles.
Factor

variahle 3 4 , 6 C"nmn,".1Iity

Rain -0.002 -0,022 -0,085 0,·3% 0.087 -0.046 0.191

Time of",;n -(J.Ulb -0.012 -(l,U82 O,4Ul 0.035 -O.U28 0.8.15
Amount ofr.lin -0.006 0.003 -O.I·!2 O.3!11 -O.().14 -O.03B 0.597
,"octurn,,! te"'per,,'n,c 0.176 -0.010 0,021 -(Una -·().OI4 0.022 0.961
Evcning tempcr~t"rc O.W() -0.028 -O.Oli - (!.OZO -0.0,51 0.037 0.951
Latc night temperat"re 0,150 O.()O4 0.(154 -0.040 0.017 0,002 0.933
\laximum temperature 0.215 -0,030 -U,2~{j D.lll D.D.30 -D,107 0.841
Minimum temperatu,e 0.150 0.013 (W1l8 -(J.ost> 0'(J24 (LOOt; 0.928
Sundown temper.,tt,re 0.203 -0.039 -0.084 -O.WI -0.097 O,OZ7 0.8SZ
Temper.,tllre drop 0,049 0.018 -0.264 0.112 0.191 -0140 0574
;-Joctlln1;ll c10lld COver -0,027 -0.016 0.295 -0.06,5 0.045 -0.041 0.910
Evening cloud COVeT -O'()29 -O.OIS 0,283 -O.Ob9 V.OII -0,029 0.7%
Late night c1""d e()Ver -0.021 -o.on O.2Kl -0.053 0'(J71l -(UJ40 0.744
Noctuma) relative humidity -0.015 O.Z70 -0.Ob7 O.01S 0.017 0.041 (UJ57
Evening relntive humidity -0.043 O.2l:>5 -O.ll!.') -0,002 (1.037 0.0.19 O.8t;O
L~te night relative humidity O.02S 0.221 -0.120 0.07f; -0.014 0.042 0,747
P"rtime Tel~tive humidity -O,O·j() 0.224 O.Ob5 -0.022 -0.037 O.ObO (1.849
Barnn"'tric pre<.'""" -(1.016 (U02 (J.05<1 -O.ISl 0.4~9 -O.l9b 0.679
Barometric p",,,,,re change -0.082 0.032 -0,094 (UllI -O,2IS 0.142 0.493
Wind vel"eitr {).O32 -0.191 -(J.071 n,H.1> -(U)J4 O.27b 0.489
Wind directi"n -O.W6 -O.O4ll -0.022 0.113 (L'517 (1.0133 O,5(;IJ
Days since rainbll 0.022 -0.005 -0.027 0.049 0,030 -0.687 D,b05
~Ioon pha"l (J.WI O.WS 0.025 -(Wig 0.375 ()A8~ (),520

as the square of the correlation coefficient (Table 1) between that weather factor and
the activity of that species. Because the weather factors are statistically independent,
the sum of the squared correlation coefficient.<; for all significant weather factors gives
the total proportion of variation in activity explained by weather. For total rodent
activity, the estimate is the sum of the squared canonical correlation coefficients (Ta-
ble 2).

DISCUSSION
Weather has a statistically significant influence on the activity of Peromysclis
maniculatus, Clethrionomys gapped, and Napaeozapus insignis. However, the influ-
ence is small compared to total variation in activity. The amount of variation explained
by weather is slightly underestimated. Although it is unlikely that any weather hwtor
not highly correlated with those measured was omitted, some slight loss of accuracy
may have occurred if responses to weather were nonlinear. The most seriOllS example
ofthis should follow from our assumption of a linear response to time of day of rainfall.
Comparisons with the results of analysis of variance show the loss in explained vari-
ation to be minimal and about equal for all three rodents. Vickery and Bider (1978),
showed nonlinear responses to be rare for Sorex cine reus.
Hayward (1965) and McNab (1970, 1978) suggested that large animals maintain

TABLE 2.-Correl(/tiolls hetwecll rodent (lctitJity {Ind weather.

P6ro",~,,'C"" cte/Iorio,,"''''''' No/we"WI''''';
F"dor "'''''iw/II/''' /.i(/;'I'<'n (mig,,;.\'

1 0.072* 0.082* 0.121 *

2 0.00:3 O.14~)* 0.041
.3 0.023 0.172* 0.117*
4 O.ll ()* 0.106* il.2(j,5*
.5 0.02.5 -0.0,'34 -(j.OO7
6 -O.(J61 -0.117* -O.O(j8*
• p < O.lJ5.
February 1981 VICKERY AND BIDER-WEATHER AND RODENT ACTIVITY 143

TABLE 3.-Canonica[ correlations hetween the weather factors and rodent activity.
Canonical variahlc

Second

Factor
1 -0.371 0.139
2 -0.287 -0.605
3 -0,463 -OA15
4 -0.679 0.589
5 0.047 0.192
6 0..320 0.241
Species
Peromyscus maniClllattis -0.213 0.301
Clethrionomys gapperi -0.554 -0.843
NapaeoUlpus insignis -0.688 0.585
Canonical correlation 0.390 0.167

constant body temperature at less energetic expense than smaller animals. In this
study P. maniculatus was smaller (body weight = 16.7 g, SD = 4.5 g, n = 22) than
either C. gapped (body weight = 21.3 g, SD = 5.6 g, n = 26) or N. insignis (body
weight = 21.5 g, SD = 3.8 g, n = 81), but was least affected by weather. Perhaps
other physiological and anatomical adaptations, such as variations in metabolic
rate and insulation, are more important than body size among animals of similar
size. For a comparison of much greater size difference, the masked shrew, Sorex
cinereus, which is much smaller than any of these rodents, is much more influenced
by weather than any of them (Vickery and Bider, 1978).
Thibault (1969) showed that N. insignis was most active on dry nights in a moist
area with heavy ground cover near a stream. Away from the stream, N. insignis was
most active on rainy nights. He stated that N. insignis changed the area in which it
was active on rainy nights. Measurements in our study area, within 100 ill of a small
stream, may reflect spatial changes in activity of N. insignis caused by weather. Vick-
ery (1976) showed that P. maniculatus and C. gapperi also change the areas in which
they are active during rainfall. Thus, all our measurements of the influence of weather
on activity may reflect changes in both total activity and the area in which rodents are
active.
All three rodents reduced their activity on cold nights. Falls (1968) concluded that
Peromysc!ls was most active at intermediate temperatures. During this study the mean
evening temperature was 1l.9°C (SD = 4.1°C) and night temperatures were always in
or below intermediate ranges cited by Falls (1968). It is likely that the environment
was never too warm for these rodents at night. Their reaction to cold temperature is
probably an energy-conserving measure.
Falls (1953) found that P. maniculatus bairdii decreased its activity at high vapor
pressures and relative humidities. Our study showed no such response in P. m. gra-
cilis. Nocturnal relative humidity in the forest was consistently high (X = 90%, SD =
7.4%) and rarely fell below the level (75%) which Falls (1953) found an increase in
activity in the grassland subspecies. C. gapperi increased its activity at high relative
humidities. C. gapperi has a high water consumption (Getz, 1962; Brower and Cade,
1966). Decreased activity at lower humidities should minimize evaporative water loss.
Falls (1968) concluded that P. 11l. hairdii was most active at intermediate light levels
and that activity of P. m. gracilis was not affected by light. He suggested that Gentry
and Odum (1957) found captures of P. maniculatus to be correlated with cloud cover
only because cloud cover was correlated with rainfall and temperature. Our study
144 JOURNAL OF tvtAMMALOGY Vol. 62, No.1

confirmed that P. m. gracilis activity is not affected by cloud cover and suggested that
the effect of cloud cover on C. gapperi and N. insignis is independent of rainfall and
temperature. Getz (1968) found that C. gapperi was most active on cloudy, moonless
nights. Youngman (1956) found the same preference in Apodemus agraritts and Vick-
ery and Bider (1978) observed the same for Sorex cinereus. Increased activity at low
light levels may be an adaptation for avoiding predators that use visual detection of
prey.
Rainfall has a major effect on the activity of all three rodents. On rainy nights P.
maniculattls activity increases by 42% compared to nonrainy nights. Activity of C.
gapperi and N. insignis increased 69% and 51 %, respectively, during periods of rain~
fall. However, this response is variable, and explains only a small proportion of the
variability in rodent activity.
Increased small mammal activity on rainy nights has been reported many times
(Burt, 1940; Falls, 1953; Gentry and Odum, 1957; Sidorowicz, 1960; Bider, 1968; Vick-
ery and Bider, 1978), but never explained. Our study showed that the increase in
activity is independent of the benefits due to higher temperature, humidity, and more
cloud cover during rainfall. Because getting wet is not an advantage (it increases the
rate of heat loss) for a small mammal, there must be some other advantage to being
active during rainfall. The most logical possibilities are changes in food availability
and predator avoidance. The rodents that respond to rainfall include herbivores and
omnivores, so food availability is unlikely to be the cause of increased activity. Small
mammals may avoid capture by predators using acoustic and olfactory senses by being
active on rainy nights. The noise of rainfall will make small mammals more difficult
to hear and moist leaf litter will make less noise than dry leaf litter. Scent trails may
be washed away more quickly during rainfall.
As the response of small rodents to rainfall and cloud cover is more likely due to
predator avoidance than to physiological restrictions, their summer activity is nearly
independent of the restrictions imposed by weather and their small body size.

ACKNOWLEDGMENTS
We thank all of the many field assistants who helped coiled the data for this study. Drs. J. S.
Millar, R. H. Green, and L. F. Soholt, an anonymous reVIewer, and Mr. D. G. L. Innes provided
helpful comments on an earlier draft of this manuscript. This project was Stlpported hy National
Research Council of Canada operating grants to both authors.

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Department of Renewable Resources, Macdonald College, P.O. 9X lCO, Canada (present

address of Vickery: Department des Sciences biologiques, Universite du Quebec a Montreal,
C.P. 8888, Succ "A," Montreal, P.Q. H9R INS Canada). Submitted 20 July 1979. Accepted 20
April 1980.

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